Form and Motion of Donax variabilis in Flow

The coquina clam, Donax variabilis, rides flow from waves, migrating shoreward during rising tides and seaward during falling tides. This method of locomotion, swash-riding, is controlled not only behaviorally but also morphologically. The shape of this clam causes it to orient passively; a clam rotates in flow, usually in backwash, until its anterior end is upstream. Rotation is about a vertical axis through a pivotal point where the shell touches the sand. The density, weight distribution, and wedge-like shape are all important in effecting orientation. Such orientation is significant because it contributes to stability of motion. On an unoriented clam, upward lift can be higher than its underwater weight--a circumstance that results in uncontrollable tumbling. In contrast, once oriented with its anterior end upstream, a clam experiences downward lift that contributes to its stability while sliding in backwash. Furthermore, when the anterior end is upstream, drag is reduced relative to when the ventral, dorsal, or posterior ends are upstream. Since orientation occurs only above a minimum velocity, it has the effect of slowing a clam's motion over the substratum in rapid flows. Stability, drag, and speed reduction enhance a clam's ability to gain a foothold and dig in after a swashride, before wave flows can wash it off the beach and out to sea.